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Arthropod Food Webs in Arctic Tundra: Trophic Interactions And ARTHROPOD FOOD WEBS IN ARCTIC TUNDRA: TROPHIC INTERACTIONS AND RESPONSES TO GLOBAL CHANGE by ASHLEY L. ASMUS Presented to the Faculty of the Graduate School of The University of Texas at Arlington in Partial Fulfillment of the Requirements for the Degree of DOCTOR OF PHILOSOPHY THE UNIVERSITY OF TEXAS AT ARLINGTON August 2017 Copyright © by Ashley L. Asmus 2017 All Rights Reserved ii Acknowledgements First and foremost, I wish to thank my advisor, Dr. Laura Gough. She has pushed me to do my best work, and kept me sane and safe along the way. My dissertation would not have been possible without the help of many fantastic field and laboratory assistants, especially Leslie Baker, Heather Bass, Rachael Carmickle, Kathryn Daly, Taryn Flink, Fernando Hernandez, Sarah Mahon, and Thy Vo. These talented students logged dozens of hours carefully sorting insects, trudged over miles of tundra and entered thousands of data sheets. They also made my life joyful. I also want to thank my Toolik Field Station family: Natalie Boelman, John Wingfield, John Moore, Gus Shaver, Jess Gersony, Jim Laundre, Libby Fortin, Laura Van Der Pol, Kiki Contreras, Sarah Meierotto, Shannan Sweet, Jonathan Perez, Helen Chmura, Jesse Krause, Seth Beaudrault and many, many more. Linda Taylor and Sherri Echols, administrators at UTA, did everything in their power to make it easy for me to do research. Dr. Jeffrey Demuth generously opened his laboratory space to me. Dr. Majie Fan (UTA Geosciences) gave me helpful advice for the stable isotope study in Chapter 4, and Dr. Derek Sikes (University of Alaska-Fairbanks Museum of the North) has graciously agreed to safeguard my unwieldy collection of arthropods after I graduate. A collaborative grant from the National Science Foundation (NSF) provided stipend, travel, undergraduate, and research support for my work in Alaska (NSF OPP 0908602 to L. Gough). Additional field support was provided by an NSF grant for the Arctic Long-Term Ecological Research project (NSF DEB 1026843 to the Marine Biological Laboratory). Key funding for the laboratory portion of this work came from the Phi Sigma Biological Honors Society at the University of Texas at Arlington. A fellowship iii from Aarhus University facilitated my work with Toke Høye, with whom I developed the ideas for Chapter 5. Last, I want to thank my partner, Jennifer Moore. She endured four summers alone in Texas in a leaky house with sub-par air conditioning and too many animals, and still managed to tell good jokes on the phone. July 10, 2017 iv Abstract ARTHROPOD FOOD WEBS IN ARCTIC TUNDRA: TROPHIC INTERACTIONS AND RESPONSES TO GLOBAL CHANGE Ashley Asmus, Ph.D. The University of Texas at Arlington, 2017 Supervising Professor: Laura Gough Arctic ecosystems are undergoing rapid change. Terrestrial arctic arthropods (insects, spiders and others) are not only appreciably diverse, but also sensitive to their environment. As such, tundra arthropod communities and food webs could provide critical insight into the ecological consequences of global change in the Arctic. My dissertation examined the underpinnings of arthropod community and food web dynamics in arctic tundra. First, I explored how changes to plant production and plant community composition affect arthropod community composition, trophic structure and food web function. I also explored one key trophic interaction: cannibalism among wolf spiders, the most abundant terrestrial predator in most arctic systems. Last, I examined the effects of plant communities and weather on arthropod phenology and activity, key determinants of the rate and role of arthropod-mediated food web processes like predation, decomposition and pollination. Overall, my research reveals that arctic consumers are strongly limited by food availability well as weather conditions in the Arctic. Early springs, warmer temperatures, increased plant production and greater shrub dominance – key consequences of arctic global change–will affect the composition of arthropod communities and the ecological functions they perform. v Table of Contents Acknowledgements .............................................................................................................iii Abstract ............................................................................................................................... v Chapter 1 Arctic Insects as Indicators of Environmental Change, 1992-2017 ................... 1 Introduction ..................................................................................................................... 1 Concepts and Methods ................................................................................................... 2 Community Composition ............................................................................................ 2 Food Web Structure and Trophic Interactions ........................................................... 4 Phenology and Physiology ......................................................................................... 7 Authorship Information.................................................................................................... 8 Chapter 2 Long-term nutrient addition alters consumer community composition but does not increase total biomass or abundance ........................................ 9 Abstract ......................................................................................................................... 10 Introduction ................................................................................................................... 11 Materials and Methods ................................................................................................. 13 Study system ............................................................................................................ 13 Nutrient addition ....................................................................................................... 14 Arthropod sampling and processing ......................................................................... 14 Plant community response measures ...................................................................... 15 Statistical Analysis .................................................................................................... 15 Results .......................................................................................................................... 18 Plant community ....................................................................................................... 18 Arthropod abundance and biomass ......................................................................... 19 Arthropod diversity .................................................................................................... 21 Arthropod community composition ........................................................................... 21 vi Discussion .................................................................................................................... 23 Fertilization did not increase total arthropod abundance or biomass ....................... 23 Fertilization decreases plant diversity, but not arthropod richness .......................... 24 Fertilization alters both plant and arthropod community composition ...................... 25 Conclusion ................................................................................................................ 26 Chapter 3 Linear responses of food web function to a gradient of nutrient enrichment ........................................................................................................................ 34 Abstract ......................................................................................................................... 35 Introduction ................................................................................................................... 35 Methods ........................................................................................................................ 39 Study system and nutrient addition .......................................................................... 39 Arthropod community responses: community composition and biomass ................ 40 Plant community responses: biomass, ANPP, C:N and community composition .............................................................................................................. 41 Food web model ....................................................................................................... 42 Statistical analysis .................................................................................................... 44 Results .......................................................................................................................... 44 Plant community responses: biomass, C:N, ANPP and community composition .............................................................................................................. 44 Arthropod community responses: biomass and community composition ................ 45 Food web model results ........................................................................................... 46 Discussion .................................................................................................................... 48 Strikingly linear responses of insect food web, but not plants, to nutrient addition ..................................................................................................................... 48 Weak herbivore effects on
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